identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038E7929EF56F85FFF4FE6D6962CFABD.text	038E7929EF56F85FFF4FE6D6962CFABD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hybosorinae	<div><p>Hybosorinae – Clade 2:  Aporolaus +  Apalonychus</p><p>This is an exclusively Neotropical clade, with two genera:  Aporolaus and  Apalonychus (including  Apalonychus similis) (Fig.1). Arrow (1911) synonymized the monospecific genus  Aporolaus with  Dicraeodon, based on the female morphology. However, its status was revalidated by Ocampo (2010b). The present study supports the genus revalidation, since  Aporolaus is the sister-group of  Apalonychus, whereas  Dicraeodon is the sister-group of  Metachaetodus (Fig. 1).</p><p>Apalonychus is recovered as a monophyletic group with the inclusion of  Hapalonychoides (Fig.1). Martinez (1994), in the original description of  Hapalonychoides, highlighted the similarity between  Apalonychus and Hapalonychoide s, suggesting that  Hapalonychoides similis could be a female of  Apalonychus pusillus Arrow, 1911, since the male was not known at that time. This proposal is not supported here because we analysed the males of both species.</p><p>According to Martinez (1994), the external body aspects of  Hapalonychoides are the same as described for  Apalonychus, except for some differences in antennomeres of the antennal club. Nonetheless, the only difference observed in antennal club characters is the presence of setae on the proximal antennomere of the club in  Apalonychus similis . Based on the results of our hypotheses and on the lack of strong characters to separate this group into two different genera, we synonymized Hapalonychoide s with  Apalonychus .</p></div>	https://treatment.plazi.org/id/038E7929EF56F85FFF4FE6D6962CFABD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Basílio, Daniel Silva;Cherman, Mariana Alejandra;Vaz-De-Mello, Fernando Zagury;Almeida, Lúcia Massutti	Basílio, Daniel Silva, Cherman, Mariana Alejandra, Vaz-De-Mello, Fernando Zagury, Almeida, Lúcia Massutti (2023): Phylogenetic relationships in Hybosoridae (Coleoptera: Scarabaeoidea). Zoological Journal of the Linnean Society 198: 1156-1170
038E7929EF56F85FFF4FE21A95BFF9A6.text	038E7929EF56F85FFF4FE21A95BFF9A6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hybosorinae	<div><p>Hybosorinae – Clade 3:  Dicraeodon +  Metachaetodus</p><p>This is another exclusively Neotropical clade (comprised of two genera:  Dicraeodon and  Metachaetodus).  Metachaetodus is the only genus in this clade that has been included in a previous phylogeny. The study of Ocampo &amp; Hawks (2006), based on molecular data, recovered  Metachaetodus as the sister-group of  Hybosorus .</p></div>	https://treatment.plazi.org/id/038E7929EF56F85FFF4FE21A95BFF9A6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Basílio, Daniel Silva;Cherman, Mariana Alejandra;Vaz-De-Mello, Fernando Zagury;Almeida, Lúcia Massutti	Basílio, Daniel Silva, Cherman, Mariana Alejandra, Vaz-De-Mello, Fernando Zagury, Almeida, Lúcia Massutti (2023): Phylogenetic relationships in Hybosoridae (Coleoptera: Scarabaeoidea). Zoological Journal of the Linnean Society 198: 1156-1170
038E7929EF56F85EFF4FE11A9329FB50.text	038E7929EF56F85EFF4FE11A9329FB50.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hybosorinae	<div><p>Hybosorinae – Clade 4:  Kuijtenous +  Hybosorus</p><p>Two distinct genera comprised this group:  Kuijtenous (Madagascar) and  Hybosorus (including  Hybosorus alluaudi) (Afrotropical) (Fig. 1). The group has an Afrotropical and Oriental distribution, except for  Hybosorus illigeri Reiche, 1853, considered a cosmopolitan invasive species (Kuijten, 1983; Ocampo, 2002).</p><p>Inourresults,  Hybosorus isrecoveredasmonophyletic only with the inclusion of  Hybosoroides . In a revision of  Hybosorus, Kuijten (1983) remarked on the strong similarity between these genera. In that research study, the only differences presented to distinguish those two genera were the density of punctures and the presence of a smooth, median swelling on the pronotum of  Hybosoroides . Kuitjen stated that he was inclined to include  Hybosoroides in  Hybosorus, although as a subgenus. In view of the topologies presented here, the proposal of a subgenus would not be possible, but they support the synonymization of  Hybosoroides with  Hybosorus .</p><p>Hybosorinae – Clade 5:  Brenskea +</p><p>(( Hypseloderus +  Seleucosorus) +</p><p>( Frolovius +  Coilodes)) + (( Celaenochrous +  Phaeochroops) + ( Phaeochridius +  Phaeochrous))</p><p>This clade is composed of nine genera: two Palaearctic ( Brenskea and  Seleucosorus), two Neotropical ( Coilodes and  Frolovius), four Oriental ( Celaenochrous,  Hypseloderus,  Phaeochroops and  Phaeochridius) and the most diverse among  Hybosorinae genera ( Phaeochrous), which is widely distributed in the Old World and in the Australian region (Fig. 1).</p><p>Brenskea (now included in  Hybosorinae) is the sister-group of all the other genera of this branch. Nikolajev (2010) suggested that  Brenskea belongs to  Dynamopodinae ( Scarabaeidae), based on the clypeus covering the mandible. However, the present hypothesis disagrees with Nikolajev (2010), since  Brenskea has not been recovered close to  Orubesa . In addition, the topology of the clade  Brenskea +  Pachyplectrus has not been recovered, as proposed by Ocampo (2006b) and by Bai et al. (2015).</p><p>Three of the five genera used in the molecular analysis by Ocampo &amp; Hawks (2006) are grouped in this clade ( Coilodes,  Phaeochroops and  Phaeochrous). In the present hypothesis,  Phaeochroops is closer to  Phaeochrous than to  Coilodes . This is different from the hypothesis of Ocampo &amp; Hawks (2006), in which  Phaeochroops appeared as the sister-group of  Coilodes and  Phaeochrous as the sister-group of all the other  Hybosorinae tested, including  Metachaetodus and  Hybosorus .</p><p>Among the lineages recovered in this clade, one is formed by  Hypseloderus +  Seleucosorus (Fig. 1). Arrow (1909) included  Hypseloderus, hitherto monospecific, in  Troginae ( Trogidae), while Schmidt (1913) relocated it into  Hybosorinae . In the description of  Hypseloderus yupae Masumoto et al., 2013, the authors stated that the asymmetric genitalia and cupuliform antennal club would be enough to include the species within  Hybosorinae (Masumoto et al., 2013) . The present phylogenetic hypothesis (the first to include this genus) supports this statement.</p><p>The monophyly of  Coilodes (with two species included) agrees with the hypothesis proposed by Bai et al. (2015).  Frolovius, the most recently described  Hybosorinae genus is placed as the sister group of  Coilodes .</p><p>Another branch clusters two Oriental genera,  Celaenochrous +  Phaeochroops . Kuijten (1984) considered  Celaenochrous as close to  Phaeochrous . However, the angled shape of the mandible and some other homoplasies positioned this genus closer to  Phaeochroops than to  Phaeochrous (Fig. 2).</p><p>The relationship of  Phaeochridius +  Phaeochrous in the same clade is retrieved for the first time, since  Phaeochridius has not been tested in any of the previous analyses proposed for  Hybosoridae or its subfamilies (Grebennikov et al., 2004; Ocampo, 2006b; Ocampo &amp; Hawks, 2006; Ballerio &amp; Grebennikov, 2016).</p><p>OUTGROUPS:  ORUBESA</p><p>The position of  Orubesa in  Scarabaeoidea is controversial because this genus is usually placed in  Dynamopodinae ( Scarabaeidae). Some authors have included it in  Hybosoridae (Allsopp, 1984; Nikolajev, 1999), but none of them tested this hypothesis in a phylogenetic analysis. Only Ballerio &amp; Grebennikov (2016) included  Orubesa athleta (Semenov, 1895) in the analysis of  Ceratocanthinae . Similar to our results (Figs 1, 2), Ballerio &amp; Grebennikov (2016) recovered  Orubesa as more related to  Ochodaeidae than to  Hybosoridae or  Scarabaeidae (this family is represented in both studies by  Aphodiini). The difference in Ballerio &amp; Grebennikov’s (2016) hypothesis is in the position of  Orubesa as the sister-group of the clade formed by  Glaphyridae +  Ochodaeidae . Thus,  Orubesa does not belong to  Hybosoridae, but its position in  Scarabaeoidea remains questionable.</p><p>The relationships of  Thinorycter (another  Dynamopodinae genus) are also inconsistent with its position within  Scarabaeoidea . Nikolajev (1999) suggested that  Thinorycter belongs to  Hybosoridae, similarly to  Orubesa . We obtained only one specimen of  Thinorycter; its inclusion in the final analysis was not possible, due to several missing morphological structures necessary to code in the character matrix. None of the preliminary analyses placed  Thinorycter in  Hybosoridae . Recently, Li et al. (2019) has proposed a hypothesis based on morphological data, in which  Thinorycter appears as the sister-group of  Glaphyridae (represented by  Eulasia,  Glaphyrus and  Pygopleurus) and distant from  Orubesa, suggesting that this genus does not belong to  Scarabaeidae, as it has been classified, or even to  Hybosoridae, as suggested by Nikolajev (1999).</p><p>HYBOSORIDAE SISTER-GROUP</p><p>Glaphyridae + ( Ochodaeidae +  Orubesa) were recovered as the sister-group of  Hybosoridae (Figs 1, 2), agreeing with the hypothesis based on the DNA sequence of eight nuclear genes proposed by McKenna et al. (2015). Despite this result, the relationships among  Hybosoridae,  Glaphyridae and  Ochodaeidae are still uncertain. Several phylogenetic analyses, all of them with morphological and molecular characters, have yielded divergent results (Browne &amp; Scholtz, 1999; Ocampo &amp; Hawks, 2006; Smith et al., 2006; Lawrence et al., 2011; Zhang et al., 2018; McKenna, 2019).</p><p>In the phylogenetic hypothesis based on 134 morphological characters from adults and larvae proposed by Browne &amp; Scholtz (1999),  Hybosoridae appear as the sister-group of  Ochodaeidae . The hypothesis proposed by Browne &amp; Scholtz (1999) was supported in the molecular analysis proposed for  Hybosoridae by Ocampo &amp; Hawks (2006). According to the hypothesis proposed by Smith et al. (2006), based on molecular data, these three  Scarabaeoidea families are polytomic.</p><p>In more comprehensive analyses, the sister-group of  Hybosoridae is even more uncertain, as shown in the morphological hypothesis proposed by Lawrence et al. (2011) for  Coleoptera, based on 516 characters from adults and larvae, in which  Hybosoridae are the sister-group of the clade  Glaresidae + ( Glaphyridae +  Melolonthidae). In addition, in the analysis based on molecular data proposed by Zhang et al. (2018),  Hybosoridae appear as the sister-group of  Scarabaeidae, and both form a clade with  Glaphyridae .</p></div>	https://treatment.plazi.org/id/038E7929EF56F85EFF4FE11A9329FB50	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Basílio, Daniel Silva;Cherman, Mariana Alejandra;Vaz-De-Mello, Fernando Zagury;Almeida, Lúcia Massutti	Basílio, Daniel Silva, Cherman, Mariana Alejandra, Vaz-De-Mello, Fernando Zagury, Almeida, Lúcia Massutti (2023): Phylogenetic relationships in Hybosoridae (Coleoptera: Scarabaeoidea). Zoological Journal of the Linnean Society 198: 1156-1170
